Pair bond and breeding success in Blue Tits Parus caeruleus and Great Tits Parus major

Data from 939 nests of the Blue Tit Parus caeruleus and 1008 nests of the Great Tit P. major from nestboxes provided in superabundance in mixed forest study sites between 1976 and 2001 were analysed to examine the effects of mate retention on breeding success and the relationship between mate fidelity and site fidelity. Most birds retained their former partner (76% in Great Tits and 65% in Blue Tits). The probability of a pair divorcing was affected by male age in Great Tits, divorce being more likely in pairs with first‐year males. Great Tit pairs breeding together for a second season bred earlier, but had no higher breeding success than pairs breeding together for the first time. In Blue Tits laying date and start of incubation tended to be earlier in pairs breeding together for a second season, but hatching and fledging dates were not earlier than in other pairs. Great Tit pairs breeding together for two consecutive seasons bred earlier in the second season than in the first, but breeding success did not differ significantly between years. In both species, breeding performance did not differ between pairs that divorced after a season and pairs that stayed together. Thus breeding success did not determine whether a pair divorced or bred together again. Neither Blue Tits nor Great Tits improved their breeding performance through divorce. Blue Tit females even had fewer fledglings in the year after divorce than in the year before. Mate retention affected breeding site fidelity. Blue Tit females had greater breeding dispersal distances between consecutive years when re‐mating than when breeding again with the same mate. In Great Tits both males and females dispersed more when re‐mating than when retaining the former partner, suggesting that mate retention increased the chance of retaining the breeding site. In both species, breeding dispersal distances did not differ between pairs that divorced and pairs in which one mate disappeared. Because no major advantage of mate retention was evident, we suggest that mate retention evolved under different conditions than those found in study sites with high breeding densities and a superabundance of artificial nesting sites.     

THE BIOLOGY OF THE WAVED ALBATROSS DIOMEDEA IRRORATA OF HOOD ISLAND, GALAPAGOS

As a nesting species, the Waved Albatross Diomedea irrorata is restricted to Hood Island in the Galapagos archipelago where 12,000 pairs bred in 1971. Outside the islands the species occurs over the northern parts of the Humboldt Current. Two colonies were studied in detail (1970–1971). At the start of a season, males returned first to the colonies and defended a small territory. Copulation occurred without any elaborate ceremony and the female spent little time on land before laying. There was no fixed nest-site, even within a season, and birds moved their eggs considerable distances. This resulted in heavy egg losses. Younger birds bred later than older birds and laid longer but narrower eggs. The average incubation spell varied from four to five days at the extremes of the incubation period to 19 days in the middle. The average incubation and fledging periods were 60 and 167 days respectively. Pairs which lost an egg sometimes adopted the abandoned egg of another bird and successfully reared the chick. Most pairs nested in both seasons. Nesting success was extremely variable, both between years and between colonies. Between 1961 and 1971 at Punta Suarez, virtually no young were reared in four seasons. Even in 1970–71, where nesting success was good, some groups of birds deserted their eggs en masse whereas in neighbouring areas up to 80% of the pairs reared young. The main foods of the young were squid and fish. Birds did not moult wing and tail feathers at the breeding colonies, and about 50% retained some primaries for more than one season, suggesting that successful pairs had difficulty in fitting in a complete moult between breeding attempts. Old feathers were normally found among the inner primaries and at the next moult were preferentially replaced, though adjacent newer feathers were sometimes retained for another season. Some birds bred in their fourth years, but most not until a year or two older. Immatures were present at the colonies late in the breeding cycle, the youngest returning latest and remaining until the last young fledged. Survival of adults and young averaged at least 95% and 93% per annum over many years. Adults and young ringed in 1961 survived equally well. The significance of the timing of the return of immatures and of the large-scale desertion of eggs, apparently not due to food shortage or disturbance, is discussed.     

Facultative dispersal by juvenile males in the cooperative stripe-backed wren

We present genetic and demographic data documenting juveniledispersal in the cooperatively breeding stripe-backed wren (Campylorhynchusnuchalis) of Venezuela. Parentage and DNA fragment-sharing analysesrevealed 12 cases in which juveniles were unrelated to othergroup members. Of these 12 foreign juveniles, (1) all were males,(2) eight of 12 had been found with breeding pairs lacking helpersrather than with groups containing helpers, and (3) four outof seven of those observed as adults courted or sired offspringwith the dominant females in their new groups despite the strongincest avoidance of this species. Furthermore, juvenile maleshad a significant tendency to disappear from natal groups intheir first year, and singleton juveniles observed with pairsafter the breeding season were mostly males. These data supportthe hypothesis that foreign juveniles were dispersers from intactgroups and not products of conspecific brood parasitism or adoptionfollowing group dissolution. We suggest that unassisted pairsmight accept juvenile males into their groups as helpers toincrease their future reproductive success and that dispersersthemselves might leave large natal groups in which their helpingis superfluous to join small groups of nonrelatives in whichthey might soon reproduce.     

SOCIAL ORGANIZATION AND NEST-BUILDING IN THE FOREST WEAVER BIRDS OF THE GENUS MALIMBUS (PLOCEINAE)

A comparative study was made of social organization during breeding among the genus Malimbus. In M. nitens, the male chooses the nest site, builds the nest alone, guards the nest during incubation, and feeds the young; the female incubates, broods alone and with the male feeds the young. In M. malimbicus, the male chooses the nest site, builds the nest with the female and guards the nest; the female builds the nest with the male, but incubates alone. In M. racheliae and M. cassini, the nest is built by one female and a multi-male party of two or three. One male drives off the other males when the nest is completed. One male and one female incubate alternately. The female seems to be the leader of the building group, and works like a male. In M. coronatus, the nest is built by a mixed party of males and females (3–6 birds), all working together without any overt leadership. Only one male and one female however, incubate, brood and feed the young. In their morphology and behaviour, Malimbus spp. are close to the weaver birds of the genus Ploceus. M. nitens seems the least evolved species while M. cassini and M. coronatus are behaviourally the most evolved. In the last species, which has a very elaborate nest, the pair of breeding birds is helped by one to four other birds. These helpers are birds in full adult plumage, and are probably capable of breeding and may do so at another period in the long breeding season of at least six months.     

Flexible Helping Behaviour in the Azure-Winged Magpie

Helping to rear the offspring of others may be a way for younger birds to gain access to future reproduction especially when turnover of breeding opportunities is low. However, this explanation is not applicable to cases where adults also help, or when roles shift between helpers and breeders. Over a period of 6-yr, we studied a marked population of azure-winged Magpie (Cyanopica cyanus) breeding in a non-territorial, colonial system. Magpies bred in a highly flexible cooperative system, in which individuals helped at different stages of the breeding cycle, including nest building, feeding the incubating female and feeding the young and removing the faecal sacs. On average, 50% of hatched nests were assisted by helpers-at-the-nest, and nest success appeared to be positively related to the presence of helpers. Helpers were predominantly males. Although juveniles were more likely to help, both juvenile and adult birds helped. Individual birds behaved as helpers either as a first-option or after having attempted their own breeding (second-option helpers). An individual helper may assist more than one nest during the same breeding season and in different breeding seasons. Reversals between breeder and helper roles were common in both directions, within a breeding season and between years. Helping behaviour is an option for almost any member of the colony. Therefore, hypotheses related to the enhancement of future breeding opportunities for juveniles can be discarded as general explanation of helping in this species. Although the decision to help appeared to be influenced by proximal environmental conditions hindering successful breeding, the associated benefits of helping as opposed to simply recovering for future reproduction, especially for former breeders, deserve further study.     

The Social System of a Communal Breeder,the Bay-winged Cowbird Molothrus badius

Molothrus badius (bay-winged cowbird), an icterine blackbird with cooperative breeding, shares behavioural and ecological characteristics with other communal nesters: it is sedentary, has a high annual survival rate (76.2%) and a strong nest-site tenacity (mean breeding dispersal of 41.9 and 89.4 m for males and females). Behavioural data, including collective agonistic displays, suggest group territoriality. Before egg hatching most breeders occurred as single pairs showing territorial behaviour (82% of nests), and nesting was usually solitary (distances to nearest nests of 25–103 m). Most breeders were apparently monogamous, with a 2.5% incidence of extrapair copulations in the territory during clutch formation. During the nestling stage one to four helpers occurred at 95% of M. badius nests. Most helpers were 1–2 years old, but older breeding adults (mostly males) that failed to rear their own offspring helped at the end of the season. The number of helpers increased (up to 4) with nestling age. Helpers were also recruited during the postfledging period, and group size reached up to 10 adults at this stage. Helpers mobbed predators and brood parasites, and provided 35% of the nestling food. Provisioning rate was positively and significantly correlated with number of helpers, although age of nestlings was the best predictor of overall food delivery rate. The helping system was almost obligate and productivity comparisons between nests with/without helpers are not possible. Data suggest that helpers increased the breeding success per nest. The correlation between the provisioning rates of parents and helpers was negative but non-significant. In 18% of nests 3 to 4 individuals were present before the nestling period, including cases of apparently polyandrous trios and one case of joint nesting by two pairs. Within Brown 's (1987) categories of social organization M. badius is mainly group territorial with plural nesting. Habitat requirements of M. badius are wide and nest sites do not appear to limit breeding. Kinship plays a role in the social system, as 9 of 12 helpers marked as nestlings helped their parents.     

THE BIRDS OF BLYTHSWOOD AND SOME NOTES ON BIRDS OF THE DISTRICT

Wintle, C. C. &; Taylor, P. B. 1993. Sequential polyandry, behaviour and moult in captive Striped Crakes Aenigmatolimnas marginalis. Ostrich 64:115-122.

Captive Striped Crakes showed sequential polyandry, the female laying for a second male when the clutch of her first mate was about to hatch. Where aviary space permitted each male set up a breeding territory and each female defended a larger area encompassing the territories of one or two males. Non-territorial subordinate males and females did not breed. The female initiated breeding by attracting the male and soliciting copulation, and the male incubated the eggs and cared for the young. Incubation took 17–18 days, the chicks left the nest at 4–5 days of age and were fully grown and capable of flight at 46–53 days. Breeding occurred from September to March and males normally reared two broods per season. Territoriality was evident only during the breeding season. Juvenile plumage was a duller version of the sexually dimorphic adult plumage; post-juvenile moult bean at 13–15 weeks and was complete at 21 weeks. Remex moult was simultaneous and a complete moult regular1 occurred twice a year in adults, in December and April (males) and September and March/April (females).     

Family status and contributions to breeding by Florida scrub jays

Nestling care by Florida scrub jay Aphelocoma c. coerulescens breeders and helpers is quantified. Breeding males and older male helpers deliver more food than first-year helpers and older female helpers. Breeding males adjust the quantity of food they deliver inversely to that delivered by their helpers. Older female helpers, who make many nest visits without delivering food, attempt to usurp the nest-sitting role of the breeding female. The advantages of group breeding to the breeders seem clear. Pairs with helpers live longer and produce more fledglings. The advantages of being a helper, though obscure, probably include non-altruistic factors such as increased survival and, for males, increased chances of obtaining space for breeding. Food contributions of helpers seem to parallel these opportunities.     

SEX RATIOS,MORPHOLOGY AND GROWTH OF THE AFRICAN BUCK DUCK

Frost, P. G. H., Ball, I. J., Siegfried, W. R. & McKinney, F. 1979. Sex ratios, morphology and growth of the African Black Duck. Ostrich 50:220-233.

Black Ducks Anus sparsa were trapped regularly in the Eerste River Valley near Stellenbosch, South Africa. The sex ratio of adult Black Ducks did not differ significantly from parity. Males were larger and heavier than females and also had proportionately larger wing spurs which are used when fighting over mates and territories. Body mass fluctuated seasonally, being lowest during summer and highest in autumn-winter. In the southwestern Cape breeding took place from July to December after the peak of the early winter rains. Ducklings hatched when waters were dropping and there was an increase in the emergence of aquatic insects. The growth rate of ducklings in the Eerste River Valley was severely retarded compared with that of ducklings reared in captivitly. Black Ducks moulted their body feathers twice a year, the moults corresponding to the pre- and post-nuptial moults of northern hemisphere waterfowl. Moults were not accompanied by any change in plumage coloration. Body and rectrix moult took more than five weeks to complete while remex replacement required about 30 days. Males began wing moult about a month earlier than females which delayed moulting until after their young had been reared. Forty-six percent of Black Ducks trapped had noticeable plumage aberrations; individual recognition among Black Ducks appears to be an important element in their social behaviour.     

Sex-bias and timing of natal dispersal in cooperatively breeding Puff-throated Bulbuls Alophoixus pallidus

While natal dispersal can have a significant impact on population dynamics, it is typically difficult to quantify. We investigated timing of natal dispersal of the cooperatively breeding Puff-throated Bulbul Alophoixus pallidus in a tropical evergreen forest by modelling the probability of staying in or dispersing from their natal territory whilst taking into account the effects of sex, group size, and the presence of helper(s). Birds did not disperse until the beginning of and during the breeding season following the hatching year. Dispersal was strongly female-biased both in frequency and distance: most females (95%) dispersed away from their natal territories, and of those relocated, traversed 2–7 territories. In contrast, 50% of males remained in the natal territory as helpers in their second year, while relocated dispersing males crossed 1–2 territories. Natal dispersal was not influenced by either group size or the presence of helpers. Males that fledged earlier in the breeding season exhibited higher rates of philopatry than the males that fledged later, but no correlation between fledging date and philopatry was observed in females. The probability of staying in the natal territory during the second year was 0.58 ± 0.14 SE and 0.05 ± 0.04 for males and females, respectively. These findings may add to our understanding of how natal dispersal can reflect social patterns and kin structure in cooperative breeding species from a little-studied tropical forest region.     

Genetic relatedness in groups is sex-specific and declines with age of helpers in a cooperatively breeding cichlid

Kin selection can explain the evolution of cooperative breeding and the distribution of relatives within a population may influence the benefits of cooperative behaviour. We provide genetic data on relatedness in the cooperatively breeding cichlid Neolamprologus pulcher. Helper to breeder relatedness decreased steeply with increasing helper age, particularly to the breeding males. Helper to helper relatedness was age‐assortative and also declined with age. These patterns of relatedness could be attributed to territory take‐overs by outsiders when breeders had disappeared (more in breeding males), between‐group dispersal of helpers and reproductive parasitism. In six of 31 groups females inherited the breeding position of their mother or sister. These matrilines were more likely to occur in large groups. We conclude that the relative fitness benefits of helping gained through kin selection vs. those gained through direct selection depend on helper age and sex.     

Winter male plumage coloration correlates with breeding status in a cooperative breeding species

The function of colored ornaments is usually related to thesignaling of individual quality in intra- and intersexual interactions.In cooperative breeding species, where only a fraction of themale population access the breeding status and the other fractionhas the option to help breeding pairs, colored traits mightprovide the females with a reliable information on the qualityof potential mate. Males of the cooperative breeding azure-wingedmagpies (Cyanopica cyanus) display conspicuous blue plumagecoloration. Here we explored the role played by structural bluecoloration of males and the probability of becoming a breederor a helper. Birds were trapped during 4 consecutive years,and feather coloration was measured with a spectrometer. Malesthat became breeders had a more brilliant and saturated bluecoloration and showed a more violet hue in the nonbreeding periodcompared with birds that became helpers. Breeding males alsoshowed a seasonal decline in blueness, whereas the color propertiesof helpers were constant throughout the year. Blueness of individualstrapped in the nonbreeding period was positively correlatedwith body size and condition. These findings are consistentwith a scenario in which nonbreeding blue plumage colorationmay function as a signal of individual quality in the azure-wingedmagpie at the pair formation time and add to growing evidencesuggesting that the nonbreeding season appears particularlyimportant in impacting breeding roles in cooperative breedingbirds.     

THE CONSEQUENCES OF BROOD SIZE FOR BREEDING BLUE TITS. III. MEASURING THE COST OF REPRODUCTION: SURVIVAL,FUTURE FECUNDITY,AND DIFFERENTIAL DISPERSAL

To determine how the cost of reproduction varies with brood size, a population of blue tits (Parus caeruleus) breeding in Wytham Wood, England, was manipulated over a three year period. Two hundred sixteen pairs were randomly assigned 3, 6, 9, 12, or 15 nestlings; nestlings were exchanged soon after hatching. Survival of adult females (as measured by the proportion recaptured in the following winter and/or spring) declined significantly with increasing brood size in two out of three years; there was significant year-to-year variation in the relationship of recapture rate to brood size. Mean female recapture rates (averaged over the three years) declined in a linear fashion (P < 0.01). There was no significant linear or curvilinear relationship between male-recapture rate and brood size in any of the three years nor was there a significant linear or curvilinear relationship for the data averaged over the three years. Nevertheless, recapture proportions for males differed significantly with respect to brood size (χ² test, P < 0.05). The possibility that experimental brood size influences subsequent dispersal (and therefore biases measures of survival based on recapture rates to differing degrees) was examined by comparing distances moved by breeding adults from one year to the next. There was no relationship between brood size and dispersal distance within the study area for either sex, except that females given broods of three were significantly more likely to move more than 300 m than were those given broods of 6–15 young. Both males and females showed evidence of a cost with respect to future fecundity: as brood size increased, the number of surviving offspring produced in the following year decreased from 1.5–1.6 (for adults that had reared 3–6 young) to 0.4 (for those that had reared 15 young). The relationship of future reproductive success to experimental brood size did not differ among years or between the sexes. The number of eggs laid and number of young hatched in year n + 1 did not differ significantly with respect to brood size in year n; rather, differences in future fecundity reflected differences in the survival prospects of young reared in year n + 1.     

Shared parentage and incest avoidance in the cooperatively breeding acorn woodpecker

Social groups of acorn woodpeckers (Melanerpes formicivorus) range in size from unaided pairs to 15 adults. Behavioural indicators of mate guarding, assumed incest avoidance and observations of egg-laying indicate that social organization ranges from monogamous pairs to groups with up to seven male and three female putative cobreeders plus up to 10 nonbreeding helpers. In addition, groups occasionally lack a putative breeder throughout the breeding season. Here we report results from multilocus DNA fingerprinting of 372 nestlings from 123 nests in groups with putative cobreeders of one or both sexes. No extra-group fertilizations were found. Putative cobreeding males within social groups shared paternity. However, the most reproductively successful male was, on average, almost three times as successful as the next most successful and additional males only occasionally sired offspring. In contrast, cobreeding females shared parentage equally. Helpers never bred incestuously when their opposite-sex parent (or another relative, such as their uncle) held breeding status in the group. However, during breeding male vacancies, 14 nestlings were produced when helper males bred incestuously with their mother. Both male and female helpers usually became successful cobreeders with their same-sex parent following replacement of the opposite-sex breeder(s) by unrelated individuals.     

COOPERATIVE BREEDING OF THE NORTHWESTERN CROW CORVUS CAURINUS IN BRITISH COLUMBIA

The behaviour of helpers at nests of Northwestern Crows was studied on Mandarte Island and Mitlenatch Island, British Columbia. Not all nests had a helper and there was only one helper per nest. Helpers participated in varying degrees in the defence of the territory and nest, feeding of the nestlings and fledglings and they cached food on the territory. Adult males fed helpers, and helpers obtained most of their food on the adults' territory. Adults with helpers laid larger clutches and produced more fledglings per nest than adults without helpers. It is suggested that cooperative breeding in the Northwestern Crow is of recent origin.     

Genetic analysis reveals diverse kin‐directed routes to helping in the rifleman Acanthisitta chloris

The social organization of cooperatively breeding species is extremely variable, with diverse social group composition and patterns of relatedness. Species that exhibit alternative routes to helping within the same population are potentially useful systems to investigate the causes and fitness consequences of diverse evolutionary pathways to cooperative behaviour. In this study, we use microsatellite markers and field observations to describe helping behaviour and patterns of relatedness in the unusual cooperative breeding system of the rifleman Acanthisitta chloris. First, we show that rifleman helpers consist of a remarkably diverse demographic, including males and females, who may be adult or juvenile, failed breeders or nonbreeders, or even successful breeders that simultaneously feed their own brood. Adult helpers mostly helped at first‐brood nests, while first‐brood juveniles assisted their parents at second broods. Second, we show that rifleman pairs are strictly sexually monogamous, and helpers did not gain any current reproductive success through helping. Third, genotyping showed that contrary to previous assumptions, helpers were closely related to the recipients of their care and preferentially directed care towards relatives over contemporaneous nests of nonrelatives. Finally, we show that variation in helper provisioning effort was attributed to age: juvenile helpers provisioned less than adults and were less responsive to the demands of a growing brood. Overall, our results show that the diverse routes to helping in this unusual species are driven by the common theme of kinship between helper and recipients, resulting in a previously underestimated potential for helpers to gain indirect fitness benefits.     

Cichlids do not adjust reproductive skew to the availability of independent breeding options

Helpers in cooperatively breeding species forego all or partof their reproduction when remaining at home and assisting breedersto raise offspring. Different models of reproductive skew generatealternative predictions about the share of reproduction unrelatedsubordinates will get depending on the degree of ecologicalconstraints. Concession models predict a larger share when independentbreeding options are good, whereas restraint and tug-of-warmodels predict no effects on reproductive skew. We tested thesepredictions by determining the share of reproduction by unrelatedmale and female helpers in the Lake Tanganyika cichlid Neolamprologuspulcher depending on experimentally manipulated possibilitiesfor helper dispersal and independent breeding and dependingon helper size and sex. We created 32 breeding groups in thelaboratory, consisting of two breeders and two helpers each,where only the helpers had access to a nearby dispersal compartmentwith (treatment) or without (control) breeding substrate, usinga repeated measures design. We determined the paternity andmaternity of 1185 offspring from 47 broods using five to nineDNA microsatellite loci and found that: (1) helpers participatedin reproduction equally across the treatments, (2) large malehelpers were significantly more likely to reproduce than smallhelpers, and (3) male helpers engaged in significantly morereproduction than female helpers. Interestingly, in four broods,extragroup helper males had fertilized part of the brood. Nohelper evictions from the group after helper reproduction wereobserved. Our results suggest that tug-of-war models based oncompetition over reproduction within groups describe best thereproductive skew observed in our study system. Female breedersproduced larger clutches in the treatment compared to the controlsituation when the large helpers were males. This suggests thatmale breeder-male helper reproductive conflicts may be alleviatedby females producing larger clutches with helpers around.     

Advanced breeding dates in relation to recent climate warming in a Mediterranean montane population of Blue Tits Cyanistes caeruleus

Trends in the onset of breeding, clutch size and numbers of hatchlings and fledglings are examined for a Mediterranean montane population of Blue Tits (Cyanistes caeruleus) subject to recent warming in springtime monitored during 20 years. Blue Tits advanced their breeding dates in relation to mean air temperatures in April and, as a consequence, laid larger clutches. However, increases in the numbers of hatchlings and fledged young over time were not statistically significant after accounting for variables of influence. The entire breeding season seems to have been displaced towards earlier dates by adjusting breeding time to increased temperatures in prebreeding time, to which Blue Tits have been more responsive than Pied Flycatchers (Ficedula hypoleuca) in the same area. The alternative hypothesis, that interference competition with Pied Flycatchers for nestboxes and caterpillars, the main common food base of nestlings, has been the driving force behind the advancement of laying of the Blue Tit population, was not supported. However, the significant advance of breeding dates in Blue Tits has not been sufficient to overcome the precipitous decline in reproductive fitness with the advancement of the season.     

Females increase reproductive investment in response to helper-mediated improvements in allo-feeding, nest survival, nestling provisioning and post-fledging survival in the Karoo scrub-robin Cercotrichas coryphaeus

In many cooperatively-breeding species, the presence of one or more helpers improves the reproductive performance of the breeding pair receiving help. Helper contributions can take many different forms, including allo-feeding, offspring provisioning, and offspring guarding or defence. Yet, most studies have focussed on single forms of helper contribution, particularly offspring provisioning, and few have evaluated the relative importance of a broader range of helper contributions to group reproductive performance. We examined helper contributions to multiple components of breeding performance in the Karoo scrub-robin Cercotrichas coryphaeus , a facultative cooperative breeder. We also tested a prediction of increased female investment in reproduction when helpers improve conditions for rearing young. Helpers assisted the breeding male in allo-feeding the incubating female, increasing allo-feeding rates. Greater allo-feeding correlated with greater female nest attentiveness during incubation. Nest predation was substantially lower among pairs breeding with a helper, resulting in a 74% increase in the probability of nest survival. Helper contributions to offspring provisioning increased nestling feeding rates, resulting in a reduced incidence of nestling starvation and increased nestling mass. Nestling mass had a strong, positive effect on post-fledging survival. Controlling for female age and habitat effects, annual production of fledged young was 130% greater among pairs breeding with a helper, and was influenced most strongly by helper correlates with nest survival, despite important helper effects on offspring provisioning. Females breeding with a helper increased clutch size, supporting the prediction of increased female investment in reproduction in response to helper benefits.     

Temporal changes in food resources,parental feeding and breeding success of Heron Island silvereyes,Zosterops lateralis chlorocephala

The influences of the temporal change in food supply on the parental feeding effort and breeding success of silvereyes,Zosterops lateralis chlorocephala, was investigated on Heron Island, Australia. Food supply (arthropods and figs) declined as the breeding season progressed. The parental feeding rate and growth of nestlings were lower when food supply was poor. When available, dominant pairs fed their young more figs and fewer arthropods than lower ranking pairs. Dominant pairs raised heavier young than lower ranking pairs when food supply was poor, while there were no significant differences between them when food supply was rich. When food supply was rich, pairs delivering greater amounts of arthropods reared nestlings better, whereas feeding more figs did not improve growth of nestlings. When food supply was poor, pairs spending a longer time at the nest reared nestlings better.